ATALANTA Zeitschrift der Zeitschrift der Zeitschrift der Zeitschrift der „Deutschen Forschungszentrale f„Deutschen Forschungszentrale f„Deutschen Forschungszentrale f„Deutschen Forschungszentrale für Schmetterlingswanderungen"r Schmetterlingswanderungen"r Schmetterlingswanderungen"r Schmetterlingswanderungen" herausgegeben herausgegeben herausgegeben herausgegeben

von der Gesellschaft zur Fцrderung der Erforschung vvon der Gesellschaft zur Fцrderung der Erforschung vvon der Gesellschaft zur Fцrderung der Erforschung vvon der Gesellschaft zur Fцrderung der Erforschung von Insektenon Insektenon Insektenon Insektenwanderungen e.V., Mwanderungen e.V., Mwanderungen e.V., Mwanderungen e.V., Münchennchennchennchen

22. Band, Heft 1 ISSN 0171-0079 April 1991

Wanderfalter in EuropaWanderfalter in EuropaWanderfalter in EuropaWanderfalter in Europa (Lepidoptera)

Zugleich Aufruf fZugleich Aufruf fZugleich Aufruf fZugleich Aufruf für eine internationale Zusammenarbeit an der r eine internationale Zusammenarbeit an der r eine internationale Zusammenarbeit an der r eine internationale Zusammenarbeit an der ErforschuErforschuErforschuErforschung des Wanderphng des Wanderphng des Wanderphng des Wanderphänomens bei den Insektennomens bei den Insektennomens bei den Insektennomens bei den Insekten

Appeal Appeal Appeal Appeal for for for for International Cooperation in International Cooperation in International Cooperation in International Cooperation in the the the the Research Research Research Research of the of the of the of the Migration Migration Migration Migration of Insectsof Insectsof Insectsof Insects

Invitation Invitation Invitation Invitation а la collaboration internationale а l'exploration du а la collaboration internationale а l'exploration du а la collaboration internationale а l'exploration du а la collaboration internationale а l'exploration du phйnomиne migrateur des insectesphйnomиne migrateur des insectesphйnomиne migrateur des insectesphйnomиne migrateur des insectes

Llamada para Llamada para Llamada para Llamada para una una una una cooperaciцn cooperaciцn cooperaciцn cooperaciцn international para el estudio international para el estudio international para el estudio international para el estudio del del del del fenфmeno migratorio de ciertos insectosfenфmeno migratorio de ciertos insectosfenфmeno migratorio de ciertos insectosfenфmeno migratorio de ciertos insectos

von

ULF ETTSCHBERGER, ROLF REINHARDT & HARTMUT STEINIGER illustriert von GUNNAR BREHM

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Foreword

It is now exactly twenty years ago that the leadership of the DFZS (Deutsche Forschungs-gemeinschaft für Schmetterlingswanderungen) passed from the hands of Dr. h.c. KURT HARZ, the founder, to us (EITSCHBERGER & STEINIGER). This proved to be a great challenge, because we were thrown overnight, without any experience, into this position of great responsibility. The tasks with which we were faced were many faceted, and were also extremely time consuming. As an example the coworkers at home and abroad had to be incorporated as members, so that sufficient finance was forthcoming. The Atalanta was at this time subsidized by the DFG and was published at irregular intervals. In addition an informational paper had to be developed to gain the interest of possible coworkers, to answer questions and also to set new standards. This was carried out by making the information and facts at this time clear and concise. Thus was published the Atalanta 4(3) which called for the international cooperation into the research of the migrational behaviour of insects. This was published in four languages. The time has now come, that this publication is fully exhausted and a totally new publication is necessary. The required finance was requested by me on the 11th August 1989 from the DFG - however without success. As this letter throws light on the tasks with which we are confronted, and also gives accountability, I have decided to quote it in full:

"The Atalanta 4(3) was published in February 1973 with a total circulation of 5000. It has since this time been the key publication for new members as well as on request for teachers, students and University teachers and students. From the original 5000 only 40 copies remain, so that a new publication is required in which the most up to date information is made available. Like the original Atalanta 4(3) it is to be published in four lan- guanges, and have 16 colour tables. I therefore request the sum of DM 15000,-. It is impossible for the DFZS to raise this sum of money under its own team. For one and a half decades, we have not been financed from any governmental or private sources, and we have vegetated on the minimum possible finance which allows to exist. A "begging letter" in the year of the Butterfly, 1987, to all the major companies only yielded four friendly replies. Since the founding of the DFZS in 1964, it has not only been the subject of insect migration research which has been accentuated, but we have also been active in the further education of the wider public. We have received much considerable correspondence in connection with questions on Ecology or Conservation, and many replies have been sent out. We have sent detailed information to schools, institutes and individuals. I am asking for the above finance to be considered immediatиly so that our comprehensive work can be furthered." In addition I have contacted Herr Professor Dr. B. HEYDEMANN, the present Minister of the Environment in land Schleswig-Holstein, so that we also have an opportunity of obtaining funds from other areas. Dr. HEYDEMANN is the most prominent member of the DFZS. So far I have not received a reply. Regardless of the outcome, we must start work on the new publication without delay, so that precious time is not lost. Time is also an important factor with regard to the new political order in Europe. As the borders fall, there is a good possi- bility of increasing the declining membership numbers.The reason for these declining numbers is the unfair Bundesartenschutzverordnung (BArtSchV) from 19.11.1986, which is showing negative effects in Germany. Even though these regulations prohibit the marking of protected butterfly species, certain cases are allowed through special application to the correct authority (see the list of

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protected butterfly species in BArtSchV). Passages of Ata- lanta 4(3) will be reused where no changes are necessary.

We very much hope that this book will give further impulses in this area, and that the subject into the research of migrational behaviour of insects will win new friends.

Marktleuthen 22nd April 1990

ULF EITSCHBERGER A) IntroductionA) IntroductionA) IntroductionA) Introduction

The DFZS (Deutsche Forschungszentrale für Schmetterlingswanderungen) was founded and organized in 1964 by Dr. h.c. KURT HARZ, with the aim of researching the migrational behaviour of insects. The problem of insect migration cannot be solved simply in the laboratory or at the writing desk, a network of observers - professional and amateur entomologists - has been organized in Central Europe. Their work contributes to this interesting branch of entomological research. It is only the great number of observers and their detailed informations which make it possible to get a pattern of the migration, re- migration and fluctuation of the populations of the various species.

The great wanderlust of the middle Europeans in the last decades and the many reports from southern Europeans, North Africans and the inhabitants of the Middle East have been invaluable, but there is still no permanent network of observers in these areas.ln the 40 years, since 1950, in which many valuable findings have been made by WARNECKE, WITT- STADT and HARZ, many questions still remain unanswered. From the information gathered from research institutes and groups in Europe (as the Netherlands, Belgium, Great Britain, Denmark, Finland and the Alpine Countries, especially Switzerland and somewhat in Yugoslavia) migrational patterns of specific types of butterflies have been pieced together. The "GroЯraumwanderer" ("long distance traveller") travel directly to the British Isles or over Spain, Western France or the Netherlands. France, The Rhineland with the western sector of Germany, Belgium and the Netherlands are normally reached from the Meditteranean and over the western Alps. From Algeria and Tunesia migration occurs into Southern France, Sardinia, Corsica and Northern Italy and further over the central Alps into Southern Germany. From here migration occurs into Northern Europe. The flight from the east appears to be via Turkey, the Balkans and Hungary, into the Eastern Alps and Czechoslovakia, Eastern Germany and the Baltic sea or directly north to Finland via Poland and the Baltic. Proof also exists for the migration to the British Isles via the Urals over Scandinavia.

Much less information is available about the return flights and the numbers of southwards flying Butterflies.

In the past, within the framework of the Butterfly Migration Research (Wanderfalterforschung), as well as the basic details of species and numbers, investigations into maturity of the gonads was also carried out. This together with the research into different forms found in insects, allows interesting new parallels to be drawn, which lead to a better understanding of the phenomenon of

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migration. This also leads to the successful breeding of migrational types. These are indeed the starting points where laboratory based studies will continue to answer questions on insect migration. We hope that the political changes in the Eastern European countries will have a positive effect on the tasks with which we are faced, i.e. the founding of further groups with central offices, the freedom of data movement and also observational information together with the increased freedom to travel. In the former GDR where butterfly migration registration and research were dissolved in 1964, the expectations are high that such work will be restarted. The formation of a new organization will not be necessary due to the existence of the DFZS.

The numerous all year round observations of coworkers will be coordinated by the central unit and will be interpreted by specialist workers. The results together with the works on the migrational behaviour of insects together with their biology and classification will be published. So that a wider readership is achieved, further works of a lepidopterological nature will be included in this publication. Our work, and that of the specialist workers, are both honorary. The not over-expensive printing costs of Atalanta were, up until 1975, covered by the generous support of the Deutsche Forschungsgemeinschaft (German Research Council) and the Bayerisches Ministerium für Unterricht und Kultur (Bavarian Ministry for Education and Culture).

Since 1975 no finance has been forthcoming from any governmental department, so that the DFZS had to manage with membership fees and donations alone. This often lead to delay in printing when the coffers were empty or low. So that this was not a regular occurrence the printing costs were covered by the managing commitee. As we now intend to extend the work and direction in which we are going, and also wish to publish the Atalanta on a more regular basis, we ask that you subscibe to this publication. At the same time you will become a member of the DFZS - and then you will be an ambassador for our common task which lies in the reseach into the migrational behaviour in butterflies and other insects.

B) The classification of the migratory butterfliesB) The classification of the migratory butterfliesB) The classification of the migratory butterfliesB) The classification of the migratory butterflies In the following the migrating species of the order Lepidoptera shall be newly classified into different groups. This new classification is by no means meant to be a definite and rigid system, but is the best system, at least in our opinion. Since the establishment of the DFZS more divisions have arisen, and just with regards this, we can see an expansion of our knowledge which has come from many years of research. These will not be stated here (EITSCHBERGER &

STEINIGER, 1973; 1980). The phenomenon of migration shows that a visible reaction is occurring within the insects and that this cannot be separated from other biological processes. We see migration (wandern) today as the opposite to staying (bleiben). A continuum of generation following generation is only possible where the yearly conditions remain fairly constant. This is only possible in very limited areas on the earth. In most places on the world's land mass there are swings in temperature, i.e. summer/winter and/or swings in water availability, i.e. dry and rainy seasons. In these areas the continuity of life is not guaranteed and during the course of evolution certain adaptibility stages have evolved. So on one side one has the stayers (Bleiber) with certain adaptability/resting stages, and on the other one has migrators (Wanderer) with its associated migratory forms. There are parallels which can be drawn between both types, and each is

21

important in the development cycles of populations. Essentially two groups can be recognized. One group reacts immediately to environmental factors, i.e. temperature change, availability of food and daylight conditions. This is the group called the consйcutives (Konsekutive). The other group is perceptive of the changes in advance, and then if at a certain point in time inhospitable conditions arise, a signal causes a stage to be produced at this point in time,even though optimal living conditions prevail. This is the group called the prospectives (Prospektive). Because of the many types at different evolutionary levels, it is not uncommon to see transitional stages or combinations of "adaptability" stages. Because of this consideration a new classification of the migratory butterflies and an extended definition of the migratory groups is necessary.

a) Prospective migratory forms I. Eumigration

Eumigrators = Seasonal migrants 1st order

Populations of butterfly species that, at specific times during the year, leave their breeding grounds and travel to specific areas (seasonal areas). Here young are produced, an^ 'hey or their offspring migrate back to the original breeding areas to reproduce. These offspring which migrate further than the specific seasonal areas are not capable of migrating back and perish. These are called "Irrgäste".

II. Paramigration

Paramigrators = Seasonal migrants 2nd order

These are populations of species which at specific time during the year leave their breeding ground and travel to areas in which they can survive, i.e. in which they can overwinter or oversummer. After the end of the resting phase, the same individuals migrate back to the breeding areas to breed. I.e. breeding occurs in the original areas from which they come.

b) Consecutive migratory forms

III. Emigration

Emigrants = Binnenwanderer

Species which migrate within their area of occurrence and do not return to the original areas from which they came, i.e. they stay in their destined area. Those individuals which are found in distant, atypical regions are called "Irrgäste". IV. IV. IV. IV. Dismigration (Evasion)Dismigration (Evasion)Dismigration (Evasion)Dismigration (Evasion)

Dismigrators = "Sammelgruppe"

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Those species which are suspected of being migratory, area expanders (Arealerweiterer), tend towards population swings and hence population spread, tend to be members of this group. The behaviour tends to be of an irregular nature. The breeding areas are left, due to various factors, and another area is reached without any particular aim (i.e. aimless).

Proof, Explanation and Examples

Groups I and II

In the case of the seasonal migrants (Saisonwanderer) the phenomenon of migration is periodically recurrent. Another similarity is that almost the whole population leaves the area of origin.

The main difference between Seasonal migrators (Saisonwanderer) orders 1 and 2 (Eumigranten/Paramigranten) is that the first group, Seasonal migrators 1st order search for more suitable areas when unfavourable conditions arise. This is because existence is not possible through change of form. Breeding ensues immediately in the new areas. A standstill in the maturity of the gonads does not occur, so that a generation of the others will follow (e.g. Cynthia cardui). A return migration to the areas of origin by the majority of the new generations, of most species, is improbable. The possible explanation for this is that the individuals that migrate northwards meet and breed, however they originate from different areas (have different origins) and therefore different genetic backgrounds. Because of these differences, the release triggers for the biological chain reaction leading to migration do not exist or exist to a different extent. Thus a threshold may not be reached and migration does not ensue in all offspring. The observed differences in yearly migration could be based on these genetic differences.

In the seasonal migrants 2nd order the maturity of the gonads goes through a resting phase in the intermediate stages (in the dormancy period). The full maturity of the gonads occurs when the dormancy period has passed. At the same time the return migration of all individuals occurs back to the regions of origin. Thus these that emigrate and then undergo the resting phase, immigrate back. Examples are Danaus plexippus (URQUHART & URQUHART, 1976), Euplagia quadripunctaria (ELGER, 1969) and Agrotis infusa (COMMON, 1954). During the resting phase the egg cells are not fully developed, and a larger amount of fat reserve is present as an energy storage (COMMON, 1954; EITSCHBERGER, 1970). As the resting phase comes to an end, and the migratory phase starts, the fat reserves decline as the eggs mature and are laid. The amount of fat reserves present is inversely proportional to the development of the eggs and quantity laid. This is true for all lepidopteran species (EITSCHBERGER, 1970). Due to the work carried out by SCHAIDER (1976) Libythea celtis was placed on the list of migratory butterflies. These creatures migrate back from areas high in the mountains to the areas of origin. Here a further resting stage is produced in which the gonads do not show maturity. Reproduction occurs after this phase has passed. L. celtis is single brooded, whereas all other members of this group are multiple brooded.

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Due to the migratory behaviour of Danaus plexippus it is placed in the seasonal migrants 2nd order (Paramigrants). The sporadic appearance of these butterflies on the West coast of Europe, mainly in England, is probably due to "chance" transport due to highwinds (BURTON & FRENCH, 1969; WILLIAMS, 1930:225-226). URQUHART (1987) does not agree with these findings. The populations which exist on the Canary Islands and Madeira, from which one has not seen any examples of migration, could have the same origins. On these islands the ecological conditions are so ideal, during the year, that this species can produce one generation after the other without production of intermediate forms or induction of migration. Exact investigations have not yet been conducted.

Group III

The Binnenwanderer differ from the Saisonwanderer in their behaviour in that migration is neither yearly nor periodic. The potential for migration is present, and can in certain populations be induced by external factors. It is more than likely that a high population density is the decisive factor which causes migration to occur. Migration is not a prerequisite for the maintenance of populations. All Binnenwanderer are in a position to exist in their birthplace at a particular developmental stage.

Group IV

Research work into individual species of other groups is currently being decided upon.

At present classification is into groups l-IV (Species marked with an asterik * are protected by the BArtSchV)

Group I Eumigranten/Saisonwanderer 1. Ordung familia Nymphalidae

Vanessa atalanta (LINNAEUS, 1758) Cynthia cardui (LINNAEUS, 1758)

familia Sphingidae Acherontia atropos (LINNAEUS, 1758) Agrius convolvuli (LINNAEUS, 1758)

familia otidae

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* Autographs gamma (LINNAEUS, 1758)1Scotia ipsilon (HUFNAGEL, 1766)

familia Pyralidae Nomophila noctuella ([DENIS & SCHIFFERMÜLLER], 1775)

Udea ferrugalis (HÜBNER, 1796) Palpita uniorialis (HÜBNER, 1796)

Group II Paramigranten/Saisonwanderer2. Ordnung

familia Danaidae Danaus plexippus (LINNAEUS, 1758)

familia Libytheidae *Libythea celtis (LAICHARTING, 1782)

familia Arctiidae

*Euplagia quadripunctaria (PODA, 1761 ) Group III

Emigranten/Binnenwanderer familia Pieridae

Aporia crataegi (LINNAEUS, 1758) Pieris brassicae

(LINNAEUS, 1758) Pieris napi (LINNAEUS, 1758) Pieris rapae

(LINNAEUS, 1758) Pontia dapiidice (LINNAEUS, 1758) Pontia edusa (FABRICIUS, 11ll)2*Pontia chloridice (HÜBNER, 1808) Gonepteryx rhamni (LINNAEUS, 1758) *Coiias hyaie

(LINNAEUS, 1758) * Coli as erate (ESPER, 1805)3*Colias crocea (GEOFFROY, 1785) Colotis evagore (KLUG, 1829) Catopsilia fioreiia

(FABRICIUS, 1775) familia Nymphalidae

Cynthia virginiensis (DRURY, 1773) Inachis io (LINNAEUS, 1758)

1 Populations from the Mediterranean migrate to us. This species is also present north of the Alps.

These Populations belong to the Binnenwanderer. 2 Following the examination carried out by GEIGER & SCHOLL we can divide the Pontia

dapiidice complex into the "eastern type" i.e. Pontia edusa (FABRICIUS, 1777) and the "western type" i.e. Pontia dapiidice (LINNAEUS, 1758). Evidence points to the fact that the "eastern type" is the migrationally more active of the two.

3 Colias erate which is widely distributed in eastern countries and native to Hungary, has in recent years been caught in Austria. G. ROSSLER from Wunsiedel caught a cf on the 3rd August 1990 in Austria, Burgenland, Seewinkel, Apetlon (pers. comm.).

familia otidae

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Aglais urticae (LINNAEUS, 1758) *lssoria lathonia

(LINNAEUS, 1758)

Danaus chrysippus (LINNAEUS, 1758) familia Lycaenidae

*Syntarucus pirithous (LINNAEUS, 1767) *Lampides boeticus

(LINNAEUS, 1767) *Lycaena phlaeas (LINNAEUS, 1758)

*Everes argiades (PALLAS, 1771) familia Sphingidae

Daphnis nerii (LINNAEUS, 1758) Macroglossum stellatarum (LINNAEUS, 1758) *Hyles livornica (ESPER, 1779) *Hyles euphorbiae (LINNAEUS, 1758) *Hyles gallii (ROTTEMBURG, 1775) Hippotion celerio (LINNAEUS, 1758)

familia Arctiidae Utetheisa pulchella (LINNAEUS, 1758)

familia Noctuidae Scotia spinifera (HÜBNER, 1785) Ochropleura leucogaster (FREYER, 1831) Noctua pronuba (LINNAEUS, 1758) Noctua fimbriata (SCHREBER, 1759) Peridroma saucia (HÜBNER, [1808]) Mythimna loreyi (DUPONCHEL, 1827) Mythimna unipuncta (HAWORTH, 1809) Mythimna vitellina (HÜBNER, [1808]) Phlogophora meticulosa (LINNAEUS, 1758) Luperinazollikoferi (FREYER, 1836) Celaena leucostigma (HÜBNER, [1808]) Spodoptera exigua (HÜBNER, 1808]) Heliothis viriplaca (HUFNAGEL, 1766) * Heliothis maritima bulgarica DRAUDT, 1938 Heliothis ononis ([DENIS & SCHIFFERMÜLLER], 1775) Heliothispeltigera ([DENIS & SCHIFFERMÜLLER], 1775) Heliothis armigera (HÜBNER, [1808]) Heliothis scutosa ([DENIS & SCHIFFERMÜLLER], 1775) Porphyrinia parva (HÜBNER, 1785) Acontia lucida (HUFNAGEL, 1767) Acontia luctuosa (ESPER, 1786) Nycteola asiatica (KRULIKOVSKY, 1904) Macdunnoughia confusa (STEPHENS, 1850) Ctenoplusia ni (HÜBNER, 1785) Chrysodeixis chalcytes (ESPER, 1789) Clytie illunaris (HÜBNER, 1785) Ophiusa tirhaca (CRAMER, 1780)

Grammodes bifasciata (PETAGNA, 1787) Grammodes stolida (FABRICIUS, 1775)

familia otidae

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Orthonama obstipata (FABRICIUS, 1794) Rhodometra sacraria (LINNAEUS, 1767) Cyclophora puppillaria

(HÜBNER, [1799]) familia Pyralidae

Dioryctria abietella ([DENIS & SCHIFFERMÜLLER], 1775) Ostrinia nubilalis (HÜBNER, 1796)

Pyrausta sticticalis (LINNAEUS, 1761) Actenia brunnealis (TREITSCHKE, 1829)

Dolicharthria punctalis ([DENIS & SCHIFFERMÜLLER], 1775) Uresiphita limbalis ([DENIS & SCHIFFERMÜLLER], 1775)

Aporodes floralis (HÜBNER, 1785) familia Tortricidae

Zeiraphera ratzeburgiana (SAXESEN, 1840) Zeiraphera rufimitrana

(HERRICH-SCHÄFFER, 1851) Zeiraphera isertana (FABRICIUS, 1794)

Zeiraphera griseana (HÜBNER, [1799]) familia Yponomeutidae

Yponomeuta padellus (LINNAEUS, 1758) Plutella xylostella (LINNAEUS, 1758)

Group IV Sammelgruppe

1) Species supposed to be migrating

familia Papilionidae *Papilio machaon (LINNAEUS, 1758)

familia Pieridae *Colias alfacariensis RIBBE, 1905

GonepteryxCleopatra (LINNAEUS, 1767) (Gruppe III ?) familia Nymphalidae

*Nymphalis antiopa (LINNAEUS, 1758) (Gruppe III ?) *Nymphalis polychloros

(LINNAEUS, 1758) (Gruppe III ?) *Nymphalisxanthomelas ([DENIS &

SCHIFFERMÜLLER], 1775) *Hipparchia semele (LINNAEUS, 1758) familia Sphingidae

Hyloicus pinastri (LINNAEUS, 1758) familia Noctuidae

Scotia segetum ([DENIS & SCHIFFERMÜLLER], 1775) Scotia exclamationis

(LINNAEUS, 1758) Amathes c-nigrum (LINNAEUS, 1758) (Gruppe III ?) Mythimna albipuncta ([DENIS & SCHIFFERMÜLLER], 1775) Mythimna l-album

(LINNAEUS, 1767) Mythimna littoralis (CURTIS, 1827)

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*Syngrapha interrogationis (LINNAEUS, 1758) *Catocala sponsa

(LINNAEUS, 1758) *Catocala electa (BORKHAUSEN, 1792) *Catocala fraxini (LINNAEUS, 1758)

2) Species extending their area of occurrence familia Lycaenidae

*Plebicula amanda (SCHNEIDER, 1792) familia Nymphalidae

*Polygonia c-album (LINNAEUS, 1758) familia Noctuidae

Omphaloscelis lunosa (HAWORTH, 1809) *Chrysaspidia bractea

([DENIS & SCHIFFERMÜLLER], 1775)

3) Interesting species, worth watching for familia Papilionidae

*lphiclides podalirius (LINNAEUS, 1758) familia Noctuidae

Rhyacia lucipeta ([DENIS & SCHIFFERMÜLLER], 1775) Mamestra brassicae (LINNAEUS, 1758) *Cucullia absinthii (LINNAEUS, 1761) Amphipyra pyramidea (LINNAEUS, 1758) Amphipyra berbera svenssoni FLETCHER, 1968 Hoplodrina blanda ([DENIS &

SCHIFFERMÜLLER], 1775) Caradrina clavipalpis (SCOPOLI, 1763)

C) Which species should be requested?C) Which species should be requested?C) Which species should be requested?C) Which species should be requested?

We would like as much information as possible about those species mentioned in the above lists, i.e. qualitative, quantitative, the types of flowers visited, the type of light flown towards which is important. Especially important is stationary and observational data over the long term. Notes and markings of insects help towards the establishment of migratory behaviour.

D) How do we recognize migratory butterflies?D) How do we recognize migratory butterflies?D) How do we recognize migratory butterflies?D) How do we recognize migratory butterflies? Species of butterflies which do not migrate fly in an irregular pattern in a fixed biotop, suck nutrition from flowers or land on the ground. Migratory types of butterfly exhibit totally different behaviour. One can quote as an example the Painted Lady (Cynthia cardui). It can be observed that this butterfly flies towards one at speed and then flies past and disappears in the opposite direction. Within a short period of time the second follows, then the third etc. This shows that the process of migration is taking place, because individual butterflies strictly maintain the same direction. Also indicative of migration are speed of flight and the maintenance of a constant flight. In addition to this type of migration in which individuals travel independently, i.e. in certain intervals of time and space, there are also those types of migration in

28

which whole flocks of individuals are involved. Here movement is in a cloudlike formation, away from an area in the same direction. Objects present in the flight path are flown over or around. Migration of insects taking place at night is much more difficult to pinpoint, for obvious reasons. If during consecutive moonlight nights, a rapid increase and then decrease in insect migration is noted, the probability exists that the insects are migrating.

One must say at this point, that a certain amount of experience is required to differentiate between these butterflies which are migratory and those which are just flight active but nonmigratory. These difficulties are especially senn in the Binnenwnderer. To make a decision in how far the butterflies are faithful to their location, usage of marking is in- dispenible.

We can distinguish between active and passive migration as follows

1) Under passive migration we understand a "freighting" of insects with the aid of air currents. No attempt is made to resist this even though they have little ability to do so. The force of migration in this group is activated by external factors (EITSCHBERGER, 1970:76).

2) In contrast to passive migration, active migration is understood to be a "directed" flight, the release of which is caused by more than one component (i.e. ecological, climatic or that of a genetic nature). The destination or target area is prefixed and this also means that the butterflies may have to fly against the wind.

E) What should be undertaken so that our work is constructive?E) What should be undertaken so that our work is constructive?E) What should be undertaken so that our work is constructive?E) What should be undertaken so that our work is constructive? Our main task lies in the observation and registration of species quoted in section C). It is most important at this point to mention that not only rare species should be observed, but also the common species. Many workers have informed us that very common species e.g. P. brassicae, P. rapae or A. urticae were not registered. It is actually these species in which accurate reporting is important, because only after many years of such reporting can an accurate picture of the population dynamics be built up. The differences between the strenghts of individual populations can be harmonized with climatic as well as ecological changes. The results can possibly also be applied to less common species. So what you have to do is give the exact number or approximated number of any observed species. Statements for example like many or single are too subjective and are fairly worthless for analytical purposes.

A report is only valuable, and this is very important, if the species identity has been correctly determined, i.e. full identification has been carried out. The damage which can be done through false reporting is greater than the value of information of which one is unsure, even though it could possibly be correct. Only report those findings on which you are sure, and for which you can show personal responsibility. For this purpose, the capture of insects gives more certainty. If you are still unsure please send a specimen to the DFZS. We will send details of identification to you.

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F) What sort of important points should a report contain?F) What sort of important points should a report contain?F) What sort of important points should a report contain?F) What sort of important points should a report contain?

1) General

1 ) Place of the observation. 2) Date and period over which the area was observed. 3) Behaviour of the butterflies i.e. general flight direction, do they remain in a particular area or not. 4) The frequency expressed in numbers. 5) The behaviour between the sexes, if this can be determined.

II) Information about migratory flights (very important)

1) In which direction are the butterflies flying? 2) How many butterflies passed by in a definite time span? 3) How many individuals were involved in the flight past? 4) In which way did the butterflies fly and what was the time gap between flights? 5) For how long did the fly past of butterflies last? 6) At what height above the ground did the butterflies fly? 7) Was the flight direction maintained by all individuals? 8) Was there a pause in flight so that individuals could rest or obtain nutrition? 9) How did the butterflies tackle obstacles? Were they flown over, or around? 10) In which direction did those butterflies travel, which had been caught and then released? 11) Did the insect flight follow particular natural paths e.g. river valleys, mountain passes, forest paths,

forest edges etc.? 12) What were the condition and size of the butterflies like? 13) What relationsship could be determined between the sexes in the sample size? (It is important to

carry out an investigation into the genitalia of the three Saisonwanderers.) 14) Were the egg cells fully developed or not? Was the fat body reduced or not? 15) Was the fly past or flight of the insects accompanied by other insect types? 16) What were the weather conditions like, especially the wind velocity/direction etc.?

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Ill) Registration cards

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All information should be written on a registration/information card with the above format. These can be obtained by workers, free of charge, from the DFZS. The information should be noted as soon as possible after an excursion is completed, so that errors are avoided. Please try not to send information in the form of a letter, as far more time is required to interpret information. Transfer of information must not lead to errors. One card should be filled out for each species. More than one card should be used for one species if observations come from a different country, province, another department or governmental office. This is helpful to interpret the information on a geographical basis.

G) How are the Butterflies marked?G) How are the Butterflies marked?G) How are the Butterflies marked?G) How are the Butterflies marked?

So that a clear picture is obtained over a specific time period, the butterflies should be marked by coworkers. For this purpose self adhesive marking labels, supplied by the DFZS, are utilized. Each label consists of two pieces, on one piece is written "Send to DFZS D-8671 Marktleuthen" and on the other a registered number. These are stuck together over a hole in the front lamina of the wing. The hole is produced with a "holer" or a fine pair of nail scissors. See figure 2.

fig. 2: The attachment of the marking label to the front wing of a butterfly.

The results so far with the marking of butterflies have not been very encouraging. We are however convinced that as soon as the network of observers is enlarged and that many butterflies are marked, that this will prove successful. Please make sure that only these butterflies are marked which are caught in a particular area and are released immediately. We are not in favour of the marking of artificially bred butterflies, whose caterpillars developed under artificial conditions (in a laboratory, room etc.). These do not react in the same way as these butterflies

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whose caterpillars developed under natural conditions. Migratory behaviour is also different and can also lead to an anormally in results.

We would ask that workers carry an example of this information with them all times so that they are correctly orientated into the methodology for both common and rare species. For a cost of DM 20,- we will send a second copy to all members of the DFZS.

Translation:

BERNARD MICHAELIS

WeihererstraЯe 34 D-8650 Kulmbach

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Literatur (Auswahl)

ARNSCHEID, W. (1976): Ist Gonepteryx Cleopatra (LINNE, 1767) ein Wanderfalter? (Lep. Pieridae). -Atalanta 7:141-142.

BACK, W. (1976): Ist Pontia chloridice (HÜBNER, 1808) ein Binnenwanderer 2. Ordnung? (Lep. Pieridae). - Atalanta 7:22-24.

BURMANN, K. (1973): Wandernde Kleinschmetterlinge (Microlepidoptera). - Atalanta 4:353- 360. BURMANN, K. (1976): Wandernde Lepidopteren (Macro- und Microlepidoptera). Eine weitere Ergänzung

der Wanderfalterliste. - Atalanta 7:49-55. BURTON, J. F. & R. A. FRENCH (1969): Monarch butterflies coinciding with American passerines in Britain

and Ireland. - British Birds 62:493-494. COMMON, I. F. B. (1954): A study of the adult Bogong Moth, Agrotls infusa (BOISD.)

(Lepidoptera: Noctuidae), with special references to its behaviour during migration and aestivation. - Australian Journal of Zoology 2:223-263.

EITSCHBERGER, U. (1970): Wanderfalterbeobachtungen im Mai 1969 in Spanien. Zugleich ein Beitrag über Gonadenuntersuchungen an Wanderfaltern. - Atalanta 3:17-42.

EITSCHBERGER, U. (1972a): Eine neue Rasse von Amathes c-nigrum (LINNE) aus Spanien (Lep. Noctuidae). - Atalanta 4:14-21.

EITSCHBERGER, U. (1972b): Pieridae, Weisslinge. Im Jahresbericht 1971 der Deutschen Forschungszentrale für Schmetterlingswanderungen. - Atalanta 4:65-110.

EITSCHBERGER, U. (1976): Stand der Wanderfalterforschung - Ziele und Wege. - Unverцffentlichtes Manuskript eines Vortrages, gehalten am 13.VII.1976 am Zoologischen Institut der Universität Bonn und im Herbst 1976 am Museum Bielefeld.

EITSCHBERGER, U. & H. STEINIGER (1973): Aufruf zur internationalen Zusammenarbeit an der Erforschung des Wanderphänomens bei Insekten. - Atalanta 4:133-192.

EITSCHBERGER, U. & H. STEINIGER (1977): Zur Wanderfaltersituation im Jahr 1976 auf der Iberischen Halbinsel, Gran Canaria und Mallorca. - Atalanta 8:234.

EITSCHBERGER, U. & H. STEINIGER (1979): Aporia crataegi ein Binnenwanderer 1. Ordnung (Lep. Pieridae). - Atalanta 10:378-381.

EITSCHBERGER, U. & H. STEINIGER (1980): Neugruppierung und Einteilung der Wanderfalter für den europäischen Bereich. - Atalanta 11:254-261.

ELGER, B. (1969): Freilandstudien zur Biologie und Цkologie von Panaxia quadripunctaria (Lep., Arctiidae) auf der Insel Rhodos. - Oecologia 2:162-197.

FELTWELL, L. (1976): Migration of Hipparchia semele. -J. Res. Lepid. 15:83-91. HARZ, K. (1963): Merkblatt der DFZS über Wanderinsekten, deren Beobachtung, besonders über

Wanderfalter. - A. Hauck, Kцnigshofen. HARZ, K. & H. WITTSTADT (1957): Wanderfalter - Die Neue Brehm-Bücherei, Heft 191, Ziemsen Verlag,

Wittenberg Lutherstadt. HIGGINS, L. G. & N. D. RILEY (1978): Die Tagfalter Europas und Nordwestafrikas. - Verlag Paul Parey,

Hamburg und Berlin. KLEINDIENST, K. (1978): Arealerweiterung und Migration von Lycaeniden im Wolfsburger Raum. Beitrag

zur Kenntnis der Wolfsburger Lycaenidenfauna (Lep. Lycaenidae). -Atalanta 9:156-164. KOCH, M. (1964): Zur Gruppeneinteilung der Wanderfalter. -Z. Wien. Ent. Ges. 49:131-143.

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MARSCHNER, G. (1976): Über die Arealausbreitung von Plebicula amanda (SCHNEIDER, 1792) (Lep. Lycaenidae). - Atalanta 6:216-218.

MAZZUCCO, K. (1966): Das Totenkopfjahr 1964 (Acherontia atropos L). - Z. Wien. Ent. Ges. 51:42. MAZZUCCO, K. (1969): Bericht der Beobachtungsstation WeiЯsee 1969. - Verlautbarungen der

Naturwissenschaftlichen Arbeitsgemeinschaft am Haus der Natur, Salzburg: 18-19. REINHARDT, R. (1987): Zur Ausbreitung von Plebicula amanda SCHN. unter besonderer Berücksichtigung

der DDR-Südbezirke (Lep., Lycaenidae). - Ent. Nachr. Ber. 31:57-62. REINHARDT, R. & K. HARZ (1989): Wandernde Schwärmerarten. - Die Neue Brehm-Bücherei, Heft 596,

Ziemsen Verlag, Wittenberg Lutherstadt. RETZLAFF, H. (1979): Wandernde Eulenfalter über der Ostsee. - Atalanta 10:172-173. REZBANYAI, L. (1978): Wanderfalter in der Schweiz 1977. - Atalanta 9:305-337. SCHAIDER, P. (1976): Libythea celtis (LAICHARTING, 1782) ein Wanderfalter? (Lep. Liby- theidae). -

Atalanta 7:19-21. STEINIGER, H. (1972): Wanderfalterbeobachtungen im Herbst 1971 in Portugal. - Atalanta 4:49-53. URQUHART, F. A. & N. R. URQUHART (1976): Monarch Butterfly (Danaus plexippus) Overwintering

Population in Mexico. - Atalanta 7:56-61. URQUHART, F. A. (1987): The Monarch Butterfly: International Traveler. - Nelson-Hall-Verlag, Chicago. WARNECKE, G. (1950): Wanderfalter in Europa. - Z. Wien. Ent. Ges. 35:100-106. WILLIAMS, C. B. (1930): The Migration of Butterflies. - Oliver and Boyd, Edinburgh. WOLFSBERGER, J. (1966): Die Macrolepidopteren-Fauna des Gardaseegebietes. - Memorie del Museo

Civico di Storia Naturale Verona 12:40-41.

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